Protection Ouart9l'Iy Vol. 3(3) 1988 115

1. Aean.boce~u. pen ••gonus (L.) Britton & Rose A. pentagonus, sword pear, is native to REVIEWS southern Texas, peninsular Florida, the Keys and thence south to northern (Benson 1982). It has spiny stems upright to 2-3 m and then arching over to form dense clumps. The large white flowers open at night and ripe fruit is eaten by birds and mammals. The means of introduction to is unknown; the earliest record is from Springsure, Qld, in 1926. The species now occurs in wooded country over a large area Distribution and biological control of north and south of Gogango in Central Queensland and in several small patches species in eastern Australia between Charters Towers and Jandowae (Figure I). The mealybug Hypogeococcus festerionus (Lizer y Trelles) (Hemiptera: J. R. Hoaklng Pseudococcidae), introduced for the biolo­ New South Wales De partment 01 Agriculture, Agricultural Researc h Centre, gical control of harrisia cactus, E. martin;;, R.M.B. 944. Ta mworth , N.SW. 2340 is established at Gogango and provides reasonable control. No other insects are R. E. McFadyen recorded from this cactus. Quee nsla nd Depa rtment 01 La nds . The Ala n Fletche r Research Station. P.O. Box 36. Sherwood, Queensla.nd 4075 2. Echlnopsis muiliple. (Pfeiffer & Otto) N. D. Murray Zucco Department 01 Genetics and Human Va riation, La Trobe University, E. mUltiplex is native to Brazil (Mann Bundoora . Victoria 3083 1970). Commonly cultivated in gardens for its delicate trumpet-shaped pink flowers, it forms dense clumps and has occasionally become naturalized from garden escapes. Summary often best suited 10 certain habitats. Such Only one infestation, near Clermont in data, where known, are included here. Central Queensland, is known to exist at Maps are presented detailing present distri­ Reviews such as those of Dodd 1940, Mann present (Figure 2). butions of cactus species in eastern main­ 1969, 1970, Moran and Zimmenmann 1984 No biological control of this cactus has land Australia. Also included are the areas and Julien 1987 should be consulted for been attempted and there are no records of of origin, common Rames, plant features further information on biological control insects feeding on it in Australia. and the first recorded Australian occur­ of cactus. rence for each species. Biological control agents for each cactus are listed along with J. Erlocereus 1bonplandll (parm. ex information on whether these agents are Materials and methods Pfeiffer) Rlccob. established in Australia and the degree of Mann (1970) lists Eriocereus regelii Weng control achieved by them. Nomenclature of cactus speci es used in this as naturalized north of Jandowae, Qld. In publication corresponds to that used by 1982 an area of several hectares of an Telford (1984). Most of the species are Eriocereus sp. with spineless fruit was Introduction described in Telford (1984) and those discovered at Springton near Duaringa in species not covered by him are described Central Queensland (Figure 3). These Dodd (1940) published a distribution map in Britton and Rose (1919-1923) and for stricto (Haw.) Haw. varieties , reaching over 3 m tall when sup­ Benson (1982). ported by trees, appeared identical to E. and since then the only detailed cactus dis­ The maps were compiled from informa­ bonplandii, a common species in the tribution maps to be published were for tion on files and from offi ce rs of the Eriocereus martin;; (Labouret) Riccob. in eastern Chaco of (McFadyen Prickly-Pear Destruction Commission of 1986). When the plants at Jandowae were Queensland (Mann 1970, Johnston and New South Wa les and the Department of Lloyd 1982, McFadyen 1986) and Opunrio subsequently re-examined, they appeared to Lands in Queensland. museum specimens be the same species. However, no authori­ auran/iaea Lindley in New So uth Wales and from the personal observations of the (Hosking and Deighton 1979). This paper tative identification has yet been obtained authors. (Telford 1984). updates the earlier maps and describes New information on biological control more species than are mentioned in pre· The mealybug H. festerionus has been agents has been gathered from personal released on E. ?bonplandii at Jandowae vious publications. Superficial maps for observations and unpublished experiments. most species were provided by I. Telford, and Springton. resulting in satisfactory with assistance from the authors, for 'Flora conlrol in both areas. of Australia: Volume4'(Telford 1984). The Dlatributlon 01 cactua epeelea and their maps will be of interest to those working biological control agents 4. Erioet.. u. mlrtlnU (Labou,et) Rlccob. on biological control of weeds and weed control in general. Dried specimens of each The cactus species are listed in alphabetical E. martinii, harrisia cactus, is a native of of the cactus species discussed here are order with information on areas of origin. the Chaco of Argentina and Paraguay slored in the herbarium of the Canberra common names, first records in ~ustralia, (McFadyen 1979). Harrisia is the name Botanic Gardens. present distribution and biological control Britton and Rose adopted for the Biological control agents for each cactus agents. Some plant features are included, Eriocereus (Britton and Rose 1919-23). species are listed along with comments on particularly those relating to presence or E. martin;; grows as a rambling tangled whether they are established in Australia absence of spines, clump-forming ability mass with 1 or 2 central spines (10-35 mm and the desree of control achieved by them. and whether plants are dispersed by long) and 5 to 7 radial spines (I~ mm These insects, mites and pathogens 3re or vegetatively. long) per areole. This species has large 116 Planl PrOieclion Ouarterly Vol. 3(3) 1988

Acanthocereus pentagonus + Echinopsis multiplex • Opuntia cylindrlca undatus Opunti. deject. Nyctocereus serpentlnus 6 Opuntia e/atior

Se/enlcereus macdona/d/ae • Opuntia ficus-indica

o Opuntia microdasys

cugo Opuntia paraguayensis km Townsville CIDI v Opuntia robusta t T ownSVi ll e~([])Opuntia streptacantha

0100 ~

Longreach + ?: ------.--- - .---.... ------___:.t'..._': _ 9£~hampton ~ ----- . ~ Roma .... . • Brisbane ~ ______- G-O-O-n-d-i-w~in~

Figure 1 (above) Distribution of pentagonus. Hylocer· o eus undarus. Nycrocereus serpentinus and Selenicereus macdonald/ae. Shaded areas are over , Tamworth. 100 ha. other areas are less than 100 ha .

Figure 2 (right) Distribution of Echinopsis multiplex. Opuntia cylin· • Sydney drica. O. dejecta, O. elatior. O. ficus·indica, O. C~nberra. microdasys, O. paraguayensis, O. rabusta, 0. strep­ * Opuntia sulphurea tacantha, 0. sufphurea, 0. tunicata and O. sp. . . 'joconoxtle'. Shaded areas are over 100 ha , other Opuntia tunicata areas are less than 100 ha. * o Opuntia sp. 'joconoxtle'

white Howers which open at night. E. mar­ Curculionidae), also became established osus but causes little damage. A . cereicola tin;; is mainly spread by seed, fruits being (McFadyen and Tomley 1981) but the latter was released but did not establish. No other eaten by both birds and mammals. has since died out (McFadyen 1986). Caeto­ in sects have been recorded on E_ tortuosus The earliest record of in troduction of blastis new sp. (Lepidoptera: Pyralidae) in Australia. E. martinii is as a pot plant brought to (McFadyen 1980) was released but has not Gatton in about 1886 (Mann 1967). E. mar­ become established on E, martinii. Other 6. Hylocereus uod.lus (Haw.) Britton linii was first recorded as a weed in insects found feeding on E, martinii are & Rose Queensland in 1935 (McFadyen and li sted in Table I. Tomley 1981). This species now occurs in H. undatus, moonli ght cactu s, is a native discontinuous areas from the Collinsv ille of tropical America (Telford 1984). It is a district in central Queensland to nonhern 5. Eriocereus tortuosus (James Forbes ex common epiphyte in gardens throughout New South Wales south of Goondiwindi Otto & A. Dietr.) Riccob. Queensland, growing up to 30 m high on (Figure 3). E. lorluosus is a native of Argentina trees, panicularly iron barks. Aerial roots E. martinii has been the object of a (Telford 1984). This cactus is also known attach the stems closely to the bark. Dur­ biological control program conducted for as harrisia cactus in Australia. E. lorluosus in g the summer there is profuse blooming the Queensland Deparlment of Lands by grows as a rambling tangled mass with 6 of large white fl owers opening at night, but the Commonwealth I nstitute of Biological to 10 radial spines (1-3 em long) and I to the large red fruit rarely form . Control. Insects on E. martinii were studied 3 central spines (2.5-6 em long) per areole. Small patches of H , undalUS are natural­ and tested in Argentina. Following these The flowers are white. . ized in and near towns from Cairns to tests, insects specific to E. martinii and E. torluosus was planted in a garden at Brisbane (Figure I). Individual infestations related genera were sent to Queensland and Western Creek Homestead, Millmerran, increase quite rapidly but as fruit are rare, released by the Department of Lands in Qld, in about 1888 (Mann 1970). The spread is min imal and H. undatus has never Queensland. The most success ful biologi­ largest infestation of E. lorluosus now become a problem. cal control agent introduced was H . Jesleri­ occurs in this area, although some plan1s No biological control has been attempted anus. Queensland now reli es on this insect have escaped from gardens in the Hunter against this cactus and' no insects are for control of E. martinii. H , /eslerianus Valley, New South Wales (Figure 3). recorded feeding on it. is still bei ng hand distributed to spread this H . Jesterianus causes severe damage to insect throughout E, marlinii infesta­ E. tortuosus in Queensland (McFadyen 7. Nyclocereus serpentinus (Lagasca & tions (McFadyen and Tomley 1981~ Two 1979) and is now relied on for control of Rodrigues) Brilton & Rose beetles, Alcidion cereicola Fisher (Cole­ this cactus. Caclob/aslis new sp. introduced optera: Cerambycidae) and Eriocereo­ from Argentina for the control of E. mar­ N. serpenl;nus, snake cactus, is native to phaga humeridens O'Brien (Coleoptera: tinii has possi bly established on E. lorlu- Mexico (Telford 1984). It has stems covered Plant Protection Quarterly Vol. 3(3) 1988 117

Table 1 Summary of published information on biological control agents of minor courses in north-eastern New South Wales importance not otherwise dealt with in the text and south-eastern Queensland (Figure 4). In 1933 Dac/y/opius austrinus De Lotto Cactus species Biological control ag_nt Reference (Hemiptera: Dactylopiidae), under the name D. sp. nr. con/usus (Cockerell), was Eriocereus martinii Chelinidea tabulata (Burmeister) Mann 1970 released for control of O. aurantiaca (Hemiptera: Coreidae) (Mann 1970). This insect causes a lot of Dip/acaspis echinocacti (Bouche) Mann 1970 damage particularly in hot dry conditions (Hemiptera: Diaspididae) (Hosking and Deighton 198 1, Hosking Tucumania tapiaco/a Dyar Mann 1970 1984). D. austrinus controls O. aurantiaca (Lepidoptera: Pyralidae) in Queensland, although some manual Opuntia aurantiaca Moneilema ulkei Horn Mann 1%9 redistri bution of this insect is occasionally (Coleoptera: Cerambycidae) necessary. D. austrinus is assuming an D. echinocacti Mann 1%9 in creasingly important role in control of Mimorista pu/chella/is Dyar Moran 1981 O. aurantiaca in New South Wales. Two (Lepidoptera: Pyralidae) moths, Cactob/as/is cactorum (Berg) and Tucumania tapiaco/a Dyar (Lepidoptera: Opuntia ficus-indica Archlagocheirus /unestrus Thompson Annecke and Pyralidae), also damage O. aurantiaca in (Coleoptera: Cerambycidae) Moran 1978 Australia. T. /apiaco/a was specifically Metamasius spinolae (Gyllenhal) Annecke and released for control of O. aurantiaca in (Coleoptera: Curculionidae) Moran 1978 1935, while C. cac/orum was released, Opuntia imbricata C. tabulata Anon. 1925 largely for control of O. stricta varieties, T. tapiacola Hoffman and in 1926 (Mann 1970). C. caclarum may Moran 1977 cause considerable damage to clumps of O. aurantioca but usually fails to kill all Opuntia streptacantha M. ulkei Mann 1970 joints within clumps. T. tapiaco/a most ly O/ycella junctolineella (Hulst) Mann 1970 causes minor damage to joints around the (Lepidoptera: Pyralidae) base of plants. Aphis armoraciae Cowen Opuntia stricta Eriophyes Von Siebold sp. Mann 1970 (Hemiptera: Aphididae) has been found on (Acarina: Eriophyidae) roots of O. aurantiaca (unpubl. data) but Tetranychus opuntiae Banks Mann 1970 the degree of damage caused by this aphid (Acarina: Tetranchidae) is unknown. In recent years Heliothis punc­ M . spinolae (Gyllenhal) Mann 1970 tigera Wallengren (Lepidoptera: Noctu­ idae), a native insect, has been recorded M . u/kei Horn Mann 1970 feeding on fl owers and young joints but 1970 Moneilema vario/are Thomsom Mann damage is minimal. Other insects wh ich (Coleoptera: Cerambyc idae) feed on O. ouronliaco are listed in Table Aphis armoraciae Cowen Eastop 1966 I. Two plant pathogens, Colletotrichum sp. (Hemiptera: Aphididae) close to C. capsici (Syd.) Butl. & Bisby C. tabulata Mann 1970 (Coelomycetes) and Phoma close to P. Chelinidea vittiger Uhler Mann 1970 exigua Desm. (Coelomycetes), cause minor (Hemiptera: Coreidae) damage to O. aurontiaca. This damage con/usus (Cockerell) Mann 1970 appears to be greatest in wet cool areas (Hemiptera: Dactylopiidae) fr om late winter to early summer (unpubl. D. echinocacti Mann 1970 data). Melitara prodenialis Walker Mann 1970 (Lepidoptera: Pyralidae) 9. Opunti. cylindric. (Lam.) DC. O. j untiolineella Mann 1970 O. cylindrica is native to Peru and Ecuador T. tapiaco/a Mann 1970 (Telford 1984), and is recorded by Telford Opuntia tomentosa T. opuntiae Mann 1970 (1 984) as occasionally naturalized in New D. echinocacti Mann 1970 South Wales and Vi ctoria as a garden escape. This species has been recorded rrom alongsid e the Werribee Ri ver in Victoria. A cactus which may be this species occurs nat uralized in a small area nea r Longreach, with soft spines. The night-blooming in South Africa. O. aurantiaca is th e major Qld (Figure 2). fl owers are large and white and the ripe cactus pest in New South Wales and South fruit are red. It is frequently cultivated and Africa. O. aurantiaca is also common in /0. Opunlla d_j«ta Salm-Dyck has become naturalized outside towns south-eastern Queensland where it is not throughout Queensland from Townsville to now regarded as a seri ous problem (Mann O. dejecta is probably native to Panama the New South Wales bord er (Figure I). t 970) . O. aurantiaca grows as a low sh rub (Mann 1970). It has long narrow pads with Occasional large infestations have devel­ rarely exceeding 40 cm in height , except long spines and grows as a much-b ranched , oped but it is not considered a problem. where supported by other vegetation. slightly drooping shrub. It is not known No biological co ntrol has been attempted Areoles have 3 to 7 spines up t0 4 cm long. when it was introduced into Australia but againt N . serpentinus but the mealybug This cactus is mainly spread by Hood­ a few plants are now naturalized just north H. jes/erianus attacks this species and could waters.·The Howers are ye llow, not orange of Rockhampton (Figure 2) . be used for biological control. as the species name suggests. Although O. dejecta is attacked by C. cac/orum fruit are common, of O. aurantiaca and by the cochineal Dactylopius oppunliae are not viable (Moran and Annecke 1979). Cockerell (Hemiptera: Dactylopiidae). 8. Opuntia .uranliaca Lindley The means of introduction o f O. auron­ O. aurantiaca is a native of Uruguay and tiaca to Australia is not known. The pest II. Opunlla _Iallor Miller the Entre Rios region of Argentina (Moran potential of O. aurantiaca in Australia was and Annecke 1979). This species is known recognized as early as 1911 (Maiden 19110). O. e/a/ior is a native of Panama and north­ as tiger pear in Australia and jointed cactus O. aurantioca is now common along water- ern South America (Telford 1984). This 118 Plant Protection Quarterly Vol . 3(3) 1988

~ • Eriocereus 1bonpl.ndll

e') - Eriocereus mar1inll 'q .. Opuntl•• ur.nrlac.

([b .. Erlocereus tor1uosus

~o km 0100 k;;;

...... ··e,;eriid.··· O'.q. ~ ~~~!,~.~Pt on

eRoma +

Goondiwin i •• \ \ ~. T;'mw o' th • Dubbo ...... 'ti Muswellbrook ....

Sydney

Figure 4 Distribution of Opuntia auranriaca. Shaded areas are over 100 ha, other areas are less than 100 ha. The shaded area for Queensland is the approximate distribution. As for New Flgure 3 Distribution of Eriocereus ?bonplandii, E. martinii and E. tor· South Wales, O. auranriaca probably follows floodplains with tuOSUS. Shaded areas are over 100 ha, other areas are less other isolated areas resulting from movement 01 joints by than 100 ha. animals and vehicles.

plant is not common in Australia (Figure able in form. Fruit, particularly of the 1964). Other insects which feed on O. ficus­ 2) . O. e/alior grows as a shrub or tree to spineless form, has been used as a food indica are li sted in Table 1. 5 m with 2 to 6 spines (rarely I) up to since prehistoric times (Benson 1982). This cactus should nOl become a major 4 cm long per areole. The fl owers are Trade has probably resulted in the current problem in Australia as the biological orange-pink. wide distribution of O. fiCU S-indica in control agents which keep it in check in Early records show that O. e/atior was tropical America. Plants grow to 7 m tall. other countries are also present in established at Liverpool, Scone, Camden Joints are largely spineless in th e va rieties Australia. and Muswellbrook in New South Wales in present in Australi a. Flowers are yellow. 1887 (Mann 1970). Maiden (1912a) said O. ficus-indica is grown throughout that this plant, recorded as Opuntia nigri­ Australia for its fruit. Spineless varieties of cans Haworth, was common at Gungai by O. ficus-indica are also grown in planta­ 13. Opuntis humi!us. (RaJ.) RaJ. 1912 and plants can still be found in this tions for drought fodder in South Africa O. humi/usa is a nalive of th e eastern half locality. The plant is nOt common now and and the U.S.A. There have been some of the U.S.A. (Telford 1984). This species those that have been seen were being fed escapes from cultivation in Australia is known as creepin g pear in Austra li a and on by C. cac/orum and D. opuntiae. Both (Figure 2) but these seem to be mainly eastern prickly pear in the U.S.A. In the these insects appear to cause considerable spineless varieties. The spin y form of U.S.A. this species has a wide distribution damage. Dodd (1940) also records that O. jicus·;ndica was the major cactus pest and grows under a wid e range of climates D. opun/iae was particularly effective in South Africa and Hawaii prior to bio­ varying from hot (humid to dry) summers against O. eta/ior and was used for control logical control of this species (Annecke and 10 severe winters (Benson 1982). O. hum;­ of this species in Sri Lanka (formerly Moran 1978, Holloway 1964). fuso is a low clump or mat. forming cact us. Ceylon) and India. Many species of insects have been Areoles are generally spineless , although recorded feeding on O. ficus-indica in so me may have single spines up to 5 em South Africa (Annecke and Moran 1978). long. The fl owers are yellow. O. humlfuso 12. Opuntl. ficus-indica (L.) Miller D. opuntiae, aided by hand-fell ing of hybridizes with O. stricto varieties in the O. ficus-indica, Indian fig, is probably plants, contributed most ( 0 the biological U.S.A. and apparent hybrids occur in Ihe native to i\ kxico where there are numerous control of O. ficus-indica in that country. Hunter Valley (Telford 1984) . cultivars and many hybrids with other C. cac/arum damages small plants or scal­ O. humljuso, recorded as O. opuntio species (Benson 1982). There are two types tered, succulent, terminal joints of lower (Linnaeus) Britton and Rose, wa s said 10 of plants common in cultivation, one spine· branches of large plants. In Hawaii be established in scattered quanlit y near less and the other spiny (formerly known D. opuntiae and C. caclOrurn were respon· Si ngleton by 1940 (Dodd 1940). Today as O. megacantha Salm-Dyck), both vari- sible for control of this cactus (Holloway O. humifuso occurs in Ihe same area Plant Protection Quarterly Vol. 3(3) 1988 119 and has not been reported elsewhere in for food, protection of villages and After initial successes insect numbers Australia (Figure 5). ceremonial purposes (Benson 1982). decreased following death of smaller plants There are no published reports of insects The earliest record of O. imbricato in and spraying was reintroduced in an feeding on O. humifusa in Australia. Australia was published in 1911 (Maiden attempt to eradicate this cactus. D. tomen­ Both C. cactorum and T. tapiacola have 191Ib). At this time O. imbrieata was losus was released in many areas within been found feeding on O. humifusa in grown in gardens in New South Wales and New South Wales and Queensland prior to the Hunter Valley. C. cactorum appears to was growing wild at a number of places in spraying and still persists in many of these cause the most damage and may be respon­ the Hunter Valley and north of Bathurst areas. Cryp/olaemus monlrouz;eri Mulsant sible for reducing O. humifusa to scattered along the Turon River. Today the largest (Coleoptera: Coccinellidae) sometimes clumps which now occur over its range in concentrations of rope pear are around old prevents population build-up of D. lomen­ the Hunter Valley. mining towns such as Broken Hill. Cobar tosus. Experiments conducted by Zimmer­ Hot dry summers appear to cause death. and Nymagee. Scattered plants also occur mann (pers. comm.) in South Africa have through desiccation. of many young joints. along a number of watercourses inland shown that felling of large plants, once from the Great Dividing Range to the south D. tomenlosus is established. improves the of Mt Isa, Qld (Figure 6). level of control achieved by this insect. 14. Opunti. imbrlca'. (Haw.) DC. Doctylopius tomen/osus (Lamark) is the Small plants are more susceptible to D. O. imbricota is a native of the U.S.A. insect which causes the most damage to lomentosus and these do not need to be cut and Mexico. occurring between central O. imbrico/a in Australia. Other insects and down. This control technique has also been Colorado and central Mexico (Benson mites also feed on O. imbrica/a. some are successfully applied in New South Wales 1982). This species is known as devil's rope of minor importance in biological control over recent years. Cutting down of old or rope pear in Australia. tree cholla or (Table I) but most of these have not been plants was found to be unnecessary in more coyonostole in the U.S.A. and imbricate released, or failed to become established. arid areas of New South Wales such as cactus in South Africa. O. imbricata grows in Australia (Mann 1969). Broken Hill and Co bar. to 3 m high as a small tree or thicket­ D. tomen/osus. recorded as D. newsleadi forming shrub. Two to 30 spines up to 3 cm Cockerell, was introduced to Australia in long arise from areoles and these spines are 1924 and released in 1925 (Dodd 1940). IS. Opuntla microdasys (Lehm.) Pfeiffer covered by a detachable sheath. The Howers Initially most of the smaller O. imbricato O. m;crodasys. golden bristle, is a native are purple and fruit vary from green to plants were killed and the larger plants of northern Mexico (Mann 1970). This yellow. American Indians used this cactus reduced to a few woody stems (Dodd 1940). plant is popular with growers of succulents.

•• Opuntl. hum/Ius. Opuntia Imbrlca'a

Opunt/. strict. var. dlllenii Q Over 100 he e;!). Opunt;. strict. var. strict. Less than 100 he [J).. Opuntl. strict. var. between

dlllenil and strict. 0100 ~ km 0.290 km

______.... ______Rockhampton Emerald ------

+ +

Roma. + + + Brisbane + Bourke ... Goondi"':in ++ + + + ~nver • Broken Hill + ~;t + • •• ~ :I- + .; • Tamworth + -Broken HiII-.. • !Oubba.

Sydney + Canberra.

Figure 5 Distribution of Opuntia humifusa, O. stricta var. dillenii, O. stricta var. stricta and 0. sin'cta var. between dillenii and stricta. Shaded areas are over 100 ha, other areas are less than 100 ha. All small areas of O. stricta are var . stricta with the exception of areas near Singleton, which are O. humi­ FIgure 8 Distribution of Opuntia imbricata. All lines other than Stale fusa, and south-east of Canberra which are 0. stricta var. boundaries are areas along watercourses where O. imbri­ between di/fenii and stn·cta . cata may be found. 120 Plant Protection Quarterl y VOl . 3(3) 1988

O. microdasys grows as a spreading shrub O. streptocontho was first planted at wealth Prickly Pear Board' to control all up to 60 cm high. The plant is without Gracemere near Rockhampton about 1880 problem cacti in Australia. Most of the spines but has large numbers of glochids and is now known in Queensland as West­ Board's work was directed toward control at each area Ie. The flowers are yellow wood or Gracemere pear. It is found in of O. stricta varieties by insects. to pink. wooded country over that part of Central C. cactorum was released for control of The earliest record is for a clump in the Queensland east of the Dawson range, O. stricto varieties in 1926 and is still the Pilliga Scrub in 1910 (Maiden 1914). This from the Mackenzie river north of Dingo major insect keeping these varieties under species does not appear to spread rapidly to near Wandoan, and to Gracemere and control (Monro 1967 , Myers et a/. 1981). and all known naturalizations are for small Biloela in the east (Figure 2). Its range In areas where C. cactorum cannot com­ areas (Figure 2). Mann (1969) lists O/ycella appears to be increasing but it is not con­ plete two generations per year there is poor juncrolineella (Hulst) (Lepidoptera: Pyra­ sidered a problem. control of O. stricta. At present these areas lidae) and D. opuntioe as insects which feed O. streptacantha is attacked by C. in New South Wales are mainly treated on O. microdosys but did not indicate the cactorum (Mann 1970) but damage to with herbicides. There are also occasional amount of damage caused by them. mature plants is minor. D. opuntiae is more problems with O. stricta control in sandy damaging and occurs widely on O. strepto­ coastal areas where O. stricto plants are cantha (Mann 1970). tough and dry and survival of C. caclorum 16. Opuntia paraguayensis K.Schum. larvae is poor (While 1981). D. opuntiae may be suitable for these areas. D. opun­ O. paraguayensis is a native of Paraguay 19. OpunUa strieta (Haw.) Haw. varieties and Argentina (Britton and Rose tiae was released in 1921 for control of 1919-1923). This species is known as River­ O. stricta is native to south-eastern U.S.A., O. stricta (Mann 1970). Early reports ina pear in New South Wales, after the area east coast of Mexico, northern South indicate that this insect provided control in in which it occurs. O. paraguayensis grows America, West Indies, Cuba, Bahamas and some areas; dispersal of C. coctorum was to 2 m high and is largely spineless, Bermuda (Benson 1982). The Iwo major so rapid, however, that D. opuntiae was although single spines sometimes occur at varieties are O. stricta var. stricto (= O. only a minor factor in initial control (Dodd areoles. These spines are up to 4 cm long. inermis DC.) which is known as common 1940). D. opuntiae is a successful biolog­ The flowers are orange-apricot with green pest pear, common prickly pear or corn­ ical agent in hot dry areas and has caused stigmas. man pear, and O. stricta var. dillenii (Ker considerable damage in so me cooler areas. There are no records of O. paraguayensis Gawler) L. Benson (~ 0. stricto) spiny pest Many other insect species have been introduction to Australia and this species pear. Plants intermediate in form between released for control of O. stricta varieties, was not mentioned in early Commonwealth O. stricto vaT. stricto and O. stricta vaT. while further species known to feed on Prickly Pear Board bulletins. O. paroguay­ dillenii are known to occur in the U.S.A. these cacti were not introduced as they were ensis is now established along the Murray (Benson 1982) and in the Araluen Valley not sufficiently host specific (Table I, Dodd Ri ver in New South Wales, Victoria and south-east of Canberra (Telford 1984). 1940, Mann 1969, 1970). as well as in other areas of Murray (1982) details the taxonomic H. punctigera causes the death of so me western New South Wales (Figure 2) . confusion relating to O. stricto in the flower buds and flowers, and minor damage C. coctorum feeds on O. paraguayensis Australian context. O. stricta varieties grow to joints. At one site in New South Wales and can cause considerable damage. Dacty­ as erect shrubs to 2 m in height but gener­ H. punctigera caused the loss of21 .0070 of lopius ceylonicus (Green) and D. opuntiae ally only reaching 1-1 .5 m. Joints vary one year's potential fruit (unpubl. data). also feed on O. paraguayensis. On this from mainly spineless in O. stricta var. Several plant pathogens have been host, D. ceylonicus develops more rapidly stricla to 1 J spines (I -6 em long) per areole recorded on O. stricta varieties (Mann with less first in star mortality than D. in some O. stricta var. dillenii plants. 1970). In Australia Phylosticta conca va opuntiae (P. Sullivan, pers. comm.). Flowers of both varieties and intermediates Seaver (Coelomycetes) often damages O. D. ceylonicus has caused a lot of damage between varieties are yellow. O. stricta is stricta varieties growing in central and to O. paroguoyensis where this in sect has also known to hybridize with O. humijuso northern Queensland (Mann 1970) and been introduced in New South Wales. in the U.S.A. (Benson 1982) and Australia coastal areas in New South Wales. The (Telford 1984). disease occurs each year between th e The origin of O. stricta in Australia is months of May and October (late autumn 17. Opunli. robusi. Wendt. ex Pfeiffer not known, although a plam in a pot was to spring) and is not apparent at other brought to Scone in 1839 (Mann 1970). It times. Gloeosporium lunatum Ell. and Ev. o. robusta, wheel cactus, is a nalive of has been suggested that prior to this plants (Coelomycetes) and a bacterial soft rot are central Mexico (Benso n 1982). O. robusta were growing in culti vation in Parramatta also widespread in Australia and cause grows to 3.5 m and has 1-10 spines (to (Mann 1970). From here joints or cuttings some damage (Mann 1970). G. lunatum is 4 cm long) per areole. The flowers are pale were taken to many areas of New South often associated with C. cactorom larvae. yellow. . Wales and Queensland for use as POl plants There are no records o f introduction of or hedges. By 1926, 24 million hectares this plant to Australia. In 1961 O. robusto were affected by O. stricta varieties with 20. OpunUa sulphurea G. Don ex Loudon was proclaimed as a nox ious plant for about half this area being unproductive as O. sulphurea is native to Argentina and Vi ctoria (Parsons 1973) 10 which it is a resuh of O. stricto infestations (Dodd Chile (Telford 1984) and was first grown mainly restricted (Figure 2). 1940). Although O. stricta varieties are now al Wallumbilla in Queensland in 1889 Mann (1969) li sts a Ilumbt'r of in sects, present over a much larger area (Figure 5), (Mann 1970). It is a spreading shrub reach­ and a mite, which feed on O. robusta and biological control has resulted in these ing a height of 30-40 ern. One to eight long gives some indicat ion of the amount of plants only rarely reaching problem spines arise from areoles. It is naturalized damage caused by them in Mexico. No proportions. at two localities in Queensland (Figure 2) biological control of thi s cactus has been The fir st detailed search for biological but shows no tendency to spread. attempted in Auslralia. control agents for O. stricto varieties fol­ No biological control has been attempted lowed the appointment of the Queensland against this cactus and there are no records Prickly Pear Travelling Commi ss ion in of in sects attacking it in Queensland. 18. Opuntia streptacantha Lemaire 191 2. This Commission spent 2 years O. streptacantha is a native of Mexico in vesti gating natural enemies of cacti 21. Opuntia tomentosa Salm-Dyck (Telford 19~4). It grows to 3-3.5 m tall, (Johnston and Tryon 1914). The in sects branching Irom the base and forming a that they brought back 10 Australia O. tomentosa is a native of central Mexico strong trunk. The segments have numerous included a species of Cactoblastis (probably (Telford 1984). This cactus is kn own as 20-25 mm long spines al the areoles. The not C. cactorum) which failed to survive velvet tree pear or velvety tree pear in flowers are yellow and the ripe fruit dark (McFadyen 1985). In 1919 the Australian Australia. Plants grow up to 8 m tall. red or purple. Government constituted 'The Common- Joints are tomentose and largely spineless. Plant PrOlection Quart""" VOl. 3{3) t988 121 although rarely I or 2 spines (3-25 mm long) occur at areoles. The flowers are + Opunlia lomenlosa orange. O. (omen/osa was recorded as common in Queensland and present in South Opunlia vulgaris Australia by 1912 (Maiden 1912b). At this time plants were found near Goondiwindi and Warwick and these areas are probably the source of present infestations in north ~ 0100 ~ eastern New South Wales. Today O. km /omenlosa is common from just south of Townsville to north-eastern New South Wales (Figure 7). The area alfected in Queensland has increased considerably and this is now the commonest cactus in Queensland, occasionally reaching pest proportions under trees on brigaJow soils. Many species of insects and mites have been recorded as feeding on O. lomentOSQ Longreach (Mann 1969). In Australia D. opunliae ------.-- . __ ._-_.. - causes the most damage to O. tomentoso. In areas where D. opunliae does not cause sufficient damage, felling of large plants increases the level of control achieved by this insect (Anon. 1984). C. caclorum destroys many seedlings and smaller plants but causes only minor damage to large plants. Archlagocheirus funeslus Thomp­ son has been reponed to cause consider­ able damage to O. tomenlosa, particularly Walgetto following initial releases in Queensland during 1936 (Mann 1970). This beetle has become less common in recent times (Mann Dubbo_. 1970) and is now nearly extinct. Other insects causing minor damage to O. tomenlosa in Australia are listed in Table I. Sydney

22. Opuntla tunkata (L.hm.) Link & 0110 O. lunieala is native to the U.S.A. , Mexico, Cuba, Ecuador, Peru and Chile (Benson 1982). It grows as a bush or is mat·forming. Six to 10 spines up to 5 em long arise from areoles and these spines are covered by a detachable sheath. The Howers are yellow. This species has been recorded from 5 km Figure 7 Distribution of Opuntia romentosa and O. vulgaris. Shaded areas are over 100 ha, other west of Miuyack in Victoria. areas are less than 100 ha. All lines other than State boundaries are areas along water· courses where O. vulgaris may be found .

2.1. Opuntla vulgaris Mill" the cochineal insects (Anon. 1925). Later D. opunti~e reportedly feeds on O. vul­ O. vulgaris is a native of Brazil, Paraguay, imports of O. vulgaris were made during garis (Mann 1970) but tests conducted at Uruguay and Argentina (Mann 1970). the 1800s (Anon. 1925). O. vulgaris was Tamworth failed to establish this cochineal Common names used for this cactus in reportedly in every State of Australia by species on O. vulgaris. C. cae/orum also Australia include drooping tree pear and 1913 (Maiden 1913). Today this cactus is causes minor damage to this cactus. Other smooth tree pear. For a long time this mainly distributed along coastal areas of insects feed on O. vulgaris (Mann 1969) but species has been known under the syno­ Australia but also occurs in gardens and as these have not been released, or have failed nym , O. monocanlha (Willd.) Haw. O. garden escapes elsewhere (Figure 7). to become established, in Australia. vulgaris normally grows as erect plants up D. ceylonieus, a native of Brazil, Uru­ Two plant pathogens, P. concava and to 2 m high but can reach greater heights guay and Argentina (Mann 1970), is the Collelolrichum cra.ssifJ'S (Speg.) An<, have when growing amongst other tall vegeta­ main biological control agent for O. been observed causing considerable damage tion. One or two spines occur per areole on vulgaris and has been successfully used to to O. vulgaris (P. Sullivan, pers. comm.). joints but up to 12 per areole on the main control this species in South Africa (Pettey trunk; spines can reach 10 cm in length. 1948), India and Ceylon (Johnston and The flowers are yellow with red mark ings Tryon 1914, Anon. 1925) as weU as U. Op•• da .p. on the outer petaloids. Australia (Dodd 1940). This insect is pos­ This cactus, known as joconoxtle, is Captain Anhur Phillip was probably sibly the one introduced by Captain Phillip recorded by Mann (1970) as a native of responsible for the first introduction of this to Australia. The Queensland TraveUing Mexico. Plants were grown at Glenrnore plant to Australia in 1788. The first fteet Commission sent D. ceylonicus from near Rockhampton in ISS7 (Mann 1970). picked up some cochineal infested plants, Ceylon in 1912 and again in 1913. These It is a moderately large cactus, 50-100 em thought to be O. vulgaris, from Rio de insects were successfully reared on O. vul­ high, with nearly round pads bearing short Janeiro in 1787 and brought them to garis at Dulacca, Qld, and subsequently spines. The flowers are yellow and fruit Australia where Captain Phillip hoped to released. By I92S O. vulgaris was reported pink. establish a cochineal dye industry (Mann to occur as rare plants in Queensland, This species is naturalized near Rock­ 1970). No records are available on the sub­ largely as a result of D. ceylonicus (Alex­ hampton but has shown no tendency to sequent destination of either this cactus or ander 1925). spread (Figure 2). 122 Planl PIOlechon Quartelly Vol. 3(3) 1988

15. Sele.leereus mocdonoldlo. (Hooker) Discussion Fletcher Research Station for information Britton & Rose on cactus distribution in Queensland. We Distribution maps shown here cover known also wish to thank Mr A. Maguire, Mr R. S. mocdonoldioe is native to Central infestations to the time of publication. Sippel and Mr P. Sullivan for assisting in America or possibly to Uruguay and adja­ Most cactus species in eastern Australia are preparation of the maps. cent parts of Argentina in South America now considered to be controlled by insects (Britton and Rose 1919-23). It is widely though most are still extending their range. cultivated for its 30--40 cm long white Long-term surveys of O. aurontiaca distri­ Howers, the largest Howers in the sub­ bution suggest that rate of spread has Relerence. family. It is a clambering plant with aerial declined over the past 20 years (Auld et al. roots and long thin stems with prominent 1982183). Unfortunately, detailed infonna­ Alexander, W. B. (1925). Natural enemies tubercles bearing very short spines. The ripe tion on rate of spread of other cactus of prickly pear and their introduction fruit are large and reddish, and bearing species is not available. Some species such into Australia. Commonwealth of Aus­ clusters of hairs or bristles. as O. imbricata and O. vulgaris appear to tralia. Institute of Science and Industry. S. macdonaldiae has only been recorded be spreading slowly, mainly vegetatively Bulletin No. 29. as naturalized at two localities (Figure 1) through spread of segments. Other species, Annecke, D. P., and Moran, V. C. (1978). and does not appear to spread or fruit such as E. martinii, are spreading more Critical reviews of biological pest control profusely. rapidly as a result of seed dispersal by birds. in South Africa. 2. The prickly pear, No biological control has been attempted The distribution maps indicate that Opuntiajicus·indica (L.) Miller. Journal against this cactus and no insects are cactus occurs in suitable country within the of the Entomological Society of South recorded from it in Queensland. area delimited, but the actual area now Africa. 41 , 161-88. infested by each cactus may be only a small Anon. (1925). 'The Prickly Pears Acclima­ proportion. For example much o.f the tised in Australia.' Commonwealth Survey 01 cactua Inle.tatlon8 In New brigalow country of Queensland prevIOusly Prickly Pear Board. (Government Prin­ South Wale. affected by cactus is now under cultivation ter: Sydney.) (McFadyen 1984) and the cactus is confined The maps in this publication do not indi­ Anon. (1984). Tree pears and their control. to the narrow treed strips left along roads Technical Bulletin, The Alan Fletcher cate the amount of each cactus species and bet ween properties. present in each locality. Some idea of this Research Station, Queensland Depart­ Insects were the main biological control ment of Lands. amount. in New South Wales, can be agents introduced, pathogens, at the time, obtained from estimates of the areas that Auld, B. A., Hosking, J ., and McFadyen, being difficult to identify and hence R. E. (1982183). Analysis of the spread they infest (Table 2). These figures include problems arose with host specificity tests. only properties on files of the Prickly-Pear of tiger pear and parthenium weed in The only pathogens mentioned are from Australia. Australian Weeds 2, 56-60. Destruction Commission and the area Opuntia species; very few specimens of where seed-borne cacti occur include entire Barker. J. S. F. (1982). Population gene­ these fungi have been examined in Austra­ tics of Opuntia breeding Drosophila in shaded areas on maps - and probably li a and their exact idcnlity remains to be some other areas. The area of one species, Australia. In 'Ecological Genetics and clarified (J. Walker, pers. comm.). Of the Evolution', eds. J. S. F. Barker and O. stricto var. stricta, is particularly insects released, C. cactorum was the most underestimated as files are no longer kept W. T. Starmer, pp.209-24. (Academic important, bringing the major cactus spe­ Press: Australia.) for properties which have a long history of cies, O. stricta, under control. Cochineal only minor Quantities of cactus. Figures for Benson, L. (1982). 'The Cacti of the United (Dactylopius species) were also impona~t other species would be more accurate. States and Canada.' (Stanford Univer­ in controlling many of the other Opunlla sit y Press: Stanford. California.) Queensland and Victoria do not have com­ species. Cochineal appear to cause the most parable figures for cactus infestations as Britton. N. L., and Rose. J. N. (1919-23). damage in hot dry conditions (Hosking 'The Cactaceae. Descriptions and illus­ control is carried out by a number of 1984). In recent times, H. festerianus has groups of government and semi­ trations of Plants of the Cactus Family.' been important in the control of E. martin;; The Carnegie Institute of Washington, government bodies as well as individuals. and other related cacti in Queensland. In Queensland, biological control is now Publication 248 , Volumes 1-4. Wash­ A number of insects not previously men­ ington. U.S.A. relied on for the control of most cactus tioned have been found in rotting cactus. species and chemical sprays are rarely used. Dodd, A. P. (1940). The biological cam­ There is no information on whether these paign against prickly-pear. Co mm~n­ insects assist in the spread of rot in plants. wealth Prickly Pear Board Bulletin. Two species of Camptodes (Coleoptera: (Government Printer: Brisbane.) Nitidulidae) have been found in rotting seg­ Eastop, V. F. (1966). A taxonomic study Table 2 Areas infested by cactus species ments of O. auran/iaca, O. streptacantho. of Australian Aphidoidea (Homoptera). in New South Wales (1988) O. stricta and O. tomen/osa. These were Australian Journal of Zoology 14. identified as C. scutel/atus Sturm and 399-592. Species Area of C. ornatus Lacordaire, although they may Hoffmann, J. H ., and Moran, V. C. infestalion represent two colour forms of the same (1977). Pre-release st udies on Tucumania (ha) species as both usually occur together. tapiacola Dyar (Lepidoptera: Pyralidae), Drosophila aldrich; Patterson & Crow and a potential biocontrol agent against Eriocereus martinii 2003 Drosophila buzzat;; Patterson & Wheeler jointed cactus. Journal of the Entomo­ E. tor/Uosus 31 (Diptera: Drosophilidae) are also found in logical Society of South Africa 40, Opuntia aurantiaca 199568 rotting tissue of many Opuntia species 205- 9. O. elatior 2 (Barker 1982). Drosophila mainly develop Holloway. J. K. (1964). Projects in bio­ O. ficus-indica I in rots which occur after damage caused by logical control of weeds. In 'Biological O. humifusa 3484 C. cac/orum or Dactylopius species. Control of Insect Pests and Weeds', ed. O. ;mbricata 6182 P. DeBach, pp.650-70. (Chapman and O. microdasys 1.5 Hall: London.) O. paraguoyensis 459 Hosking, J . R. (1984). The effect of tem­ O. robusta 0.5 Acknowledgment. perature on the population growth O. stricta var. stricta 881534 We wi sh to thank Mr G. Ryan and staff of potential of Dactylopius austrinus De O. stricta var. between the Prickly-Pear Destruction Commission Lotto (Homoptera: Dactylopiidae), on stricta and dillenii 1143 for information on cactus infestations in Opuntia aurantiaca Lindley. Journal of 45530 O. tomentosa New South Wales and Mr G. Donnelly. Mr the Australian Entomological Society 23, O. vulgaris 1759 A. Tomley and other staff of the Alan 133-9. Plant Protection Quarterly Vol. 3(3) 1988 123

Hosking, J . R., and Deighton, P. J. (1979). (Opuntia spp.). Bulletin of Entomo­ strategies. Proceedings of the Fifth The distribution and control of Opuntia logical Research 7S, 159-68. International Symposium on Biological aurantiaea in New South Wales. Pro­ McFadyen, R. E. (1986). Harrisia cactus Control of Weeds, 22-27 July 1980, ceedings of the Seventh Asian-Pacific - a pasture pest in Queensland. In Brisbane, Australia, ed . E. S. Delfosse, Weed Science Society Conference, 26- 'Ecology of Exotic Animals and Plants', pp.383-7. (CSIRO: Melbourne.) 30 November 1979, Sydney, eds R. W. ed. R. Kitching, pp.24 I-61. (Jacaranda­ Moran, V. C., and Annecke, D. P. (1979). Medd and B. A. Auld, pp.195-200. Wiley: Brisbane.) Critical reviews of biological pest con­ Hosking, J. R., and Deighton, P. J. (1981). McFadyen, R. E., and Tomley, A. J . trol in South Africa. 3. The jointed Biological control of moisture stressed (1981). Biological control of Harrisia cactus, Opunt;a aurant;aca Lindley. Opuntia aurant;oca using Daclylopius cactus, Eriocereus martin;;, in Queens­ fournal of the Entomological Society of aust';nus. Proceedings of the Fifth land by the mealybug Hypogeococcus South Africa 42, 299-329.

International Symposium on Biological festerianus. Proceedings of the Fifth Moran, V. C' I and Zimmennann, H. G. Control of Weeds, 22-27 July 1980, International Symposium on Biological (1984). The biological control of cactus Brisbane, Australia, ed. E. S. Delfosse, Control of Weeds, 22-27 July 1980, weeds: achievements and prospects. pp.483-7. (CSIRO: Melbourne.) Brisbane, Australia, ed. E. S. Delfosse, Biocontrol News and Information 5, Johnston, T. H., and Tryon, H. (1914). pp.589-94. (CSIRO: Melbourne.) 297-320. Report of the Prickly Pear Travelling Maiden, J. H. (19110). The prickly pears Murray, N.D. (1982). Ecology and evolu­ Commi ssion, 1st November, 1912-30th of interest to Australians. Agricultural tion of the Opuntia-Cactoblastis April, 1914. Parliamentary Paper. Gazelle of New South Wales 22,321-8. ecosystem in Australia. In 'Ecological (Government Printer: Brisbane.) Maiden, J. H. (I9I1b). The prickly pears Genetics and Evolution,' eds . 1. S. F. Johnston, W. R., and Lloyd, P. (1982). of interest to Australians. Agricultural Barker and W. T. Starmer, pp.17-30. The fight against cact i pests in Queens­ Gazelle of New South Walesll, 696-8. (Academic Press: Sydney.) land. Queensland Agricultural Journal Maiden, J. H. (19120). The prickly pears Myers, J. H " Monro. J., and Murray, N. 108, 215-21. of interest to Australians. Agricultural (1981). Egg clumping, host plant sel­ Julien, M. H. (ed.) (1987). 'Biological Gazelle of New South Wales 23, 208-10. ection and population regulation in Control of Weeds: a World Catalogue Maiden, J. H. (1912b). The prickly pears Cactob/astis cactorum (Lepidoptera). of Agents and their Target Weeds', of interest to Australians. Agricultural Oecologia (Bert) SI, 7-13. 2nd ed. (Wallingford, Oxon.: CAB Gazelle of New South Wales 23, 1027-8. Parsons, W. T. (1973). 'Noxious Weeds International.) Maiden, J. H. (1913). The prickly pears of of Victoria', p.37. (lnkata Press: McFadyen, R. E. (1979). The cactus mealy­ interest to Australians. Agricultural Melbourne.) bug Hypogeococcusfesterianus (Hem.: Gazelle of New South Wales 24, 863-6. Pettey, F. W. (1948). The biological con­ Pseud ococcidae) an agent for the bio­ Maiden, J. H. (1914). The prickly pears of trol of prickly pears in South Africa. logical control of Eriocereus martinii interest to Australians. Agricultural Union of South Africa, Department of (Cactaceae) in Australia. Entomophaga Gazette of New South Wales 25, 137-8. Agricuilure, Science Bulletin No. 271, 24, 281-7. Mann, J. (1967). 'The Harrisia Cactus.' Entomology Series No. 22. McFadyen, R. E. (1980). A Cactoblastis (Government Printer: Brisbane.) Telford, I. R. H. (1984). Cactaceae. In (Lep. : Phycitidae) for the biological con­ Mann, J. (1969). Cactus-feeding insects '', Volume 4, ed. A. S. trol of Eriocereus martin;; (Cactaceae) and mites. United States National George, pp.62-80. (Australian Govern­ in Australia. Entomophaga 2S, 37-42. Museum Bulletin No. 256. ment Publishing Service: Canberra.) McFadyen, R. E. (1984). Alien weeds. In Mann, J. (1970). 'Cacti Naturalised in White, G. G. (1981). Current status of 'The Brigalow Belt of Australia.' Royal Australia and Their Control.' (Govern­ prickly pear control by Cactob/astis Society of Queensland Symposium, 23 ment Printer: Bri sbane.) CQ(:torum in Queensland. Proceedings of October 1982, ed . A. Bailey pp.I77- Monro, J. (1967). The exploitation and the Fifth International Symposium on 182. (Department of Primary Industries: conservation of resources by populations Biological Control of Weeds, 22-27 July Brisbane.) of insects. Journal of Animal Ecology 1980, Brisbane, Australia, ed. E. S. Del­ McFadyen, R. E. (1985). Larval character­ 36, 531-47. fosse, pp.609-16. (CSIRO: Melbourne.) istics of Cactoblastis spp. (Lepidoptera: Moran, V. C. (1981). The biological con­ Pyralidae) and the selection of species trol of Opuntia aurantiaea in South for biological control of prickly pears Africa: evaluation and emerging control